Novitates Paleoentomologicae No

Novitates Paleoentomologicae No

Novitates Paleoentomologicae No. 7, pp. 1–15 13 May 2014 ȱȱȱȱȱȱEulaemaȱȱ¢ȱȱ ¡ȱȱǻ ¢DZȱǼ ȱǯȱ1 Abstract.ȱȱȱęȱȱȱȱȱ¢ȱȱ¡ȱȱȱȱȱęȬ ǯȱȱEulaema (Apeulaema) zigrasiȱǰȱ ȱǰȱȱȱęȱȬȱȱ ȱȱȱ¡ȱȱȱ ȱȱȱęȱȱȱȱǯȱȱȱ ȱȱ ȱȱȱȱȱȱȱȱĚ¢ȱȱȱȱȱ ¢ȱȱǯ ȱȱȱȱȱ¡ȱȱȱȱȱȱȱȱ ¢ȱȱ ȱȱȱ¡¢Ȭȱȱȱȱȱ ȱǻǰȱŘŖŖŚDzȱà£Ȭ ǰȱŘŖŖŝǼǯȱȱȱȱȱȱȱ ȱȱȱ ȱȱȱȱȱȱȱȱȱȱ ȱȱȱȱȱȱȱȱ ¢ȱȱȱǯȱȱȱ ȱȱȱȱȱȱ¢ȱȱȱǰȱ¢ȱȱȱȬ ȱȱȱȱȱ¢ȱȱ¢ȱȱȱǯȱȱ ȱȱȱ ȱȱȱȱȱȱȱȱ¢Ȭȱȱȱȱȱȱ ȱȱ¡ȱȱǽe.gǯǰȱȱȱȱȱMastotermesȱĴǰȱDol- ichorhinotermesȱ¢ȱǭȱǰȱȱIncisitermesȱ ȱǻǰȱŗşŜşDzȱ ȱ ǭȱǰȱŗşŞřDzȱ ȱǭȱ ǰȱŗşşŗDzȱ¢ȱǭȱǰȱŘŖŖŖDzȱȱ ǭȱ ǰȱŘŖŖŝǰȱŘŖŖŝǼǰȱȱȱȱȱProplebeiaȱȱǻȱet alǯǰȱŘŖŖŖǼǰȱ 1ȱȱȱ ȱ¢ǰȱȱȱȱȱ ¢ǰȱȱȱȱ ȱŝşȱǰȱ ȱǰȱ ȱȱŗŖŖŘŚȬśŗşŘDzȱȱȱȱ¢ǰȱȱ ¢ȱ ȱȱȱȱ¢ȱǭȱ¢ȱ¢ǰȱŗśŖŗȱȱȱȮȱȱŗŚŖǰȱ ¢ȱȱ ǰȱ ǰȱ ȱŜŜŖŚśȱǻȓǯDzȱȓǯǼǯ Copyright © M.S. Engel. ȱȱĴȬȬȱřǯŖȱȱȱǻȱȬȬȱřǯŖǼǯ ȱŘřŘşȬśŞŞŖ 2 Novitates Paleoentomologicae No. 7 bugs of the genera Leptosalda Cobben and Termitaradus Myers (Cobben, 1971; Poinar & Doyen, 1992; Grimaldi & Engel, 2008; Engel, 2009a; Poinar & Heiss, 2011; Grimaldi et al., 2013)]. The bee fauna of Dominican amber is remarkably diverse, with a seem- ingly steady accumulation of taxa every couple of years (e.g., Michez et al., 2012; Engel et al., 2012; Engel & Breitkreuz, 2013). In contrast, the number of described species of bees in Mexican amber has remained constant for 55 years. Trigona (Nogueirapis) silacea Wille (1959) was described from a small series of workers and, much like Proplebeia dominicana (Wille & Chandler) in Dominican amber (Camargo et al., 2000), the species is now known from hundreds of individuals from these deposits (Engel & Michener, 2013a). With the elevation of Nogueirapis to generic rank, the species was long known as Nogueirapis silacea and compared with those species of this genus with similar facial patterns and occurring in Mesoamerica and Central America (e.g., Wille, 1964). Ca- margo et al. (2000) discussed the relationship of Wille’s species relative to Proplebeia and considered them to belong to a single lineage, and Camargo & Pedro (2007) sub- sequently formally transferred the species to Proplebeia. Given that there is quite a bit of variation among the large series of specimens now available from various amber localities in Mexico, it was initially thought that there were more than one species in the fauna. However, for virtually every difference discernible there are variants that intergrade between the size classes and no strong evidence any longer exists for the recognition of multiple species (pers. obs.). There is considerable inherent interest in P. silacea but in the absence of additional bee lineages in the fauna, Mexican amber has not inspired as much interest among melittologists as has that of the Dominican Republic. This long stagnation has finally been broken with the recent discovery of a large orchid bee (Euglossini) in Mexican amber (Fig. 1). A single female of a species of Eulaema Lepeletier de Saint Fargeau has been discovered and this new species is described herein. Admittedly, this find leaves the diversity of Mexican amber bees still confined to the corbiculate Apinae, but the discovery of any additional lineages outside of the Meliponini are of considerable interest. Hopefully the present contribu- tion will spark a steady stream of new species from other bee tribes, subfamilies, and families much in the same way the discovery of Neocorynura electra Engel broke more than a quarter century of silence after P. dominicana for the Dominican fauna (Wille & Chandler, 1964; Engel, 1995), and ushered in a nearly 20 year span in which the num- ber and diversity of Dominican amber bees has continually grown (e.g., Rozen, 1996; Michener & Poinar, 1996; Engel, 1996, 1997, 1999a, 1999b, 1999c, 2000a, 2009b; Engel & Rightmyer, 2000; Camargo et al., 2000; Greco et al., 2011; Engel et al., 2012; Engel & Breitkreuz, 2013). Eulaema presently includes 33 nominal species ranging from northern Mexico to northern Chile and Argentina (e.g., Moure et al., 2007; Nemésio & Rasmussen, 2011; Nemésio & Ferrari, 2012), with these segregated into two subgenera – Eulaema s.str. and Apeulaema Moure (Moure, 1950, 1963, 2000; Oliveira, 2006, 2007). There are four species presently documented from Mexico – Eulaema (Eulaema) meriana (Olivier), E. (E.) luteola Moure, E. (Apeulaema) polychroma (Mocsáry), and E. (A.) cingulata (Fabri- cius) [the latter species is polytypic and perhaps represents multiple taxa (Oliveira, 2006), and the form present in Mexico has at times gone by the name E. marcii Ne- mésio: Nemésio (2009)] – and all of these are taxa with broad distributions (e.g., Moure et al., 2007; Hinojosa-Díaz et al., 2009; Hinojosa-Díaz & Engel, 2012a). Several of these widespread species pose taxonomic challenges and are in need of revision, likely with as of yet unrecognized cryptic taxa (pers. obs.). Indeed, critical and comprehensive revisions of these species are needed, as is true for many groups of native bees where 2014 Engel: An orchid bee in Mexican amber 3 Figure 1. Photograph of dorsal habitus of holotype female (JZC-MX-0023B) of Eulaema (Apeu- laema) zigrasi, new species, as preserved in Early Miocene Mexican amber. the circumscription of taxa has not been considered in a modern context (Engel, 2011; Gonzalez et al., 2013). Given the tectonic changes in this region since the Early Miocene 4 Novitates Paleoentomologicae No. 7 (e.g., Iturralde-Vinent & MacPhee, 1999), it is unlikely that the fossil species discussed herein is a close relative of any of the aforementioned living taxa today occurring in the Chiapas region. Fossil orchid bees have been previously discovered in copal and Dominican amber, the latter comprising species of Eufriesea Cockerell and Euglossa Latreille (Engel, 1999a; Hinojosa-Díaz & Engel, 2007). The new species from Mexican amber is described, figured, and compared with its living congeners. MATERIAL AND METHODS Morphological terminology used follows that of Engel (1999a, 2001a) and Michen- er (2007), while the format for the description is loosely based on previous works per- taining to Euglossini (e.g., Kimsey, 1982; Moure, 2000; Oliveira, 2006; Ayala & Engel, 2008; Hinojosa-Díaz & Engel, 2007, 2011a, 2011b, 2012b; Hinojosa-Díaz et al., 2011). Photography was done using a Canon EOS 7D digital camera attached to an Infinity K-2 long-distance microscope lens and illuminated by a Xenon flash, while an ocu- lar micrometer on an Olympus SZX-12 stereomicroscope was employed for measure- ments. Measured values should be considered somewhat approximate since an opti- mal angle was not always possible. The age, origin, and paleobiota of Mexican amber have been recently summarized by Engel (2004a) and Solórzano-Kraemer (2007), while general accounts of the fossil history of bees have most recently been provided by Engel (2004b), Ohl & Engel (2007), and Michez et al. (2012), with sundry updates by Wappler et al. (2012), Engel et al. (2012, 2013, 2014), Engel & Breitkreuz (2013), Engel & Michener (2013a, 2013b), Kotthoffet al. (2013), and Dewulf et al. (2014). The bee is preserved in a large, relatively clear piece of light yellowish-orange am- ber from the mines of Chiapas, Mexico. Originally a large, polished, spherical piece clearly meant as an objet d’art, it was cut with a water-fed trimming saw and then sev- eral flat surfaces were polished, particularly from above, behind, and lateral to the bee. Given the bee’s proximity to other surfaces, only smaller windows could be prepared from below and near the face. These cuts afforded critically important views of the bee, particularly the integument. Despite these necessary preparations, observation of the specimen remains somewhat hampered owing to the presence of several large, reflective fracture planes within the amber piece and around the bee (Figs. 1–4). SYSTEMATIC PALEONTOLOGY Genus Eulaema Lepeletier de Saint Fargeau Subgenus Apeulaema Moure Eulaema (Apeulaema) zigrasi Engel, new species ZooBank: urn:lsid:zoobank.org:act:DD572414-5F21-41DF-8E28-13EA2E888731 (Figs. 1–5) Diagnosis: The new species is characterized by the overall dark brown to black coloration of the integument, which is lacking metallic highlights, its dark fuscous pubescence (ochraceous or reddish pubescence not present anywhere), and the form of the clypeal ridges as described below. Description: ♀: Total body length 14.5 mm; forewing length 10.5 mm; hind wing length 7.9 mm. Head wider than long, length 2.93 mm, width 4.36 mm. Labrum length 1.12 mm, width 1.06 mm. Clypeus moderately protuberant, ridges weak, medial ridge extending from base about two-thirds clypeal length and then disappearing, weak 2014 Engel: An orchid bee in Mexican amber 5 Figure 2. Photograph of right lateral habitus of holotype female (JZC-MX-0023B) of Eulaema (Apeulaema) zigrasi, new species. paramedial ridges present only in apical half; upper clypeal width 0.90 mm, clypeal medial length 1.22 mm. Upper interorbital distance 2.07 mm, lower interorbital dis- tance 1.76; ocellocular distance 0.42 mm; interocellar distance 0.42 mm. Scape elon- gate; first flagellomere longer than second flagellomere; second flagellomere as long as third flagellomere; first flagellomere length 0.42 mm; second flagellomere length 0.25 mm; third flagellomere length 0.25 mm (flagellomeres somewhat compressed as preserved, so individual widths were not taken as they are certainly not indicative of the width in life). Mesoscutal length 2.61 mm; intertegular distance 2.92 mm; tegula subtriangular; mesoscutellum long, projecting over metanotum and propodeum, me- soscutellar length 1.65, basal width 1.86 mm. Mesotibia shorter than mesobasitarsus, length 2.02 mm; mesobasitarsal

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